Does sex affect both individual and collective vigilance in social mammalian herbivores: the case of the eastern grey kangaroo?

In several vertebrate taxa, males and females differ in the proportions of time they individually devote to vigilance, commonly attributed to sex differences in intra-specific competition or in absolute energy requirements. However, an effect of sex on collective vigilance is less often studied (and therefore rarely predicted), despite being relevant to any consideration of the adaptiveness of mixed- vs single-sex grouping. Controlling for group size, we studied the effect of sex on vigilance in the sexually dimorphic eastern grey kangaroo Macropus giganteus, analysing vigilance at two structural levels: individual vigilance and the group’s collective vigilance. Knowing that group members in this species tend to synchronise their bouts of vigilance, we tested (for the first time) whether sex affects the degree of synchrony between group members. We found that females were individually more vigilant than males and that their vigilance rate was unaffected by the presence of males. Collective vigilance did not differ between female-only and mixed-sex groups of the same size. Vigilance in mixed-sex groups was neither more nor less synchronous than in single-sex groups of females, and the presence of males seemed not to affect the degree of synchrony between females. Sixty-six percent of vigilant acts were unique (performed when no other kangaroo was alert), and only about one unique vigilant act in every three induced a collective wave of vigilance. The proportions of vigilant acts that were unique were 60% for females but only 46% for males. However, the sexes differed little in the rates at which their unique vigilant acts were copied. This limited study shows that the differences in vigilance between male and female kangaroos had no discernible effect upon collective vigilance.     

Does the risk of encountering lions influence African herbivore behaviour at waterholes?

A central question in the study of predator–prey relationships is to what extent prey behaviour is determined by avoidance of predators. Here, we test whether the long-term risk of encountering lions and the presence of lions in the vicinity influence the behaviour of large African herbivores at waterholes through avoidance of high-risk areas, increases in group size, changes in temporal niche or changes in the time spent in waterhole areas. In Hwange National Park, Zimbabwe, we monitored waterholes to study the behaviour of nine herbivore species under different risks of encountering lions. We radio-collared 26 lions in the study area which provided the opportunity to monitor whether lions were present during observation sessions and to map longer-term seasonal landscapes of risk of encountering lions. Our results show that the preferred prey species for lions (buffalo, kudu and giraffe) avoided risky waterholes. Group size increased as encounter risk increased for only two species (wildebeest and zebra), but this effect was not strong. Interestingly, buffalo avoided the hours of the day which are dangerous when the long-term and short-term risks of encountering lions were high, and all species showed avoidance of waterhole use at night times when lions were in the vicinity. This illustrates well how prey can make temporal adjustments to avoid dangerous periods coinciding with predator hunting. Additionally, many herbivores spent more time accessing water to drink when the long- and short-term risks of encountering lions were high, and they showed longer potential drinking time when the long-term risk of encountering lions was high, suggesting higher levels of vigilance. This study illustrates the diversity of behavioural adjustments to the risk of encountering a predator and how prey respond differently to temporal variations in this risk.     

Does cadence of Iguana iguana displays facilitate individual recognition?

Iguanid lizards communicate threat, courtship and territorial advertisement through stereotyped headbob displays. It has been hypothesized that slight interindividual differences in these displays might enhance individual recognition and maintain dominance relationships within populations, but the precise display elements responsible have not been determined. The purpose of this study was to examine if experimentally induced differences in displays would decrease recognition of individual conspecifics. Specifically, social interactions between adult male green iguanas (Iguana iguana) were examined at normothermic and hypothermic body temperatures. Whereas the cadence (rate of muscle contraction) of stereotypic displays of individual lizards was slowed at hypothermic body temperature (Q ₁₀ 1.6), the amplitude (maximal muscle contraction) of hypothermic displays remained constant relative to normothermic displays. Normothermic lizards viewing a display from a hypothermic lizard responded to that display as if it were performed at a normothermic cadence. During paired encounters between lizards (one hypothermic, one normothermic), dominant/subordinate dyads within the group were unusually maintained regardless of which individual was hypothermic. The exception occurred when dominant individuals at the lowest experimental body temperatures retreated from subordinate individuals. The results suggest that individual lizards recognized their relative social status even when cadence of displays was radically altered, indicating that cadence alone probably does not function in individual recognition. Other morphological, display elements, or scent cues may provide information useful in individual recognition.Communicated by G.M. Klump     

Does the cost of polygyny in house wrens include reduced male assistance in defending offspring?

Summary This study investigated whether reduced male aid in defending offspring potentially reduces the fitness of females choosing already-mated males in the house wren (Troglodytes aedon), a small, territorial songbird. Frozen snakes were placed at 23 nests of monogamously mated males and 12 secondary nests of bigamously mated males. All presentations were made during incubation stages of females attending focal nests. Snakes were placed at nests of secondary females when nests of their primary counterparts contained young 5–9 days old. Males are most attentive to primary nests during this period and should therefore be relatively inattentive to secondary mates and nests. Nevertheless, an equal proportion of monogamous and bigamous males discovered snakes within 15 min, and mean time to discovery, when discovery occurred, did not differ with nest status. Monogamous and bigamous males were also equally likely to attack snakes physically once discovered. Monogamous males appeared no more likely to discover snakes than bigamous males for two main reasons. First, although monogamous males were near focal nests (i.e., < 10 m) more often than bigamous males, monogamous males tended to stay out of view of nests for long periods. In contrast, bigamous males always went immediately to focal nests upon arriving in their vicinity. Second, about one-third of monogamous males in this study spent much of their time during trials at the far edges of their territories advertising for secondary mates. Our experiment suggests that reduced male aid in defending nests against small, diurnal predators probably does not contribute to the cost of polygyny in house wrens. Correspondence to: L.S. Johnson     

Does supernormal stimulus influence parental behaviour of the cuckoo's host?

The supernormal stimulus hypothesis (SSH) states that a cuckoo chick should obtain more parental care than host young by means of exaggerated sensory signals. We tested the SSH by comparing parental care by reed warblers at parasitized and non-parasitized nests. A comparison of feeding rates to parasite and host chicks of the same size showed that parasitized nests received more food than non-parasitized ones with one host chick. There was an interesting relationship between average prey length and the mass of a cuckoo chick: prey length first increased with chick mass, but decreased after the cuckoo chick outgrew the average-sized host brood (three to four young at fledging). This might be expected if fosterers reduced the selectivity of their foraging behaviour when trying to satisfy the supernormal food demands of the parasitic chick. This suggestion is supported by the finding that the relationship between nestling mass and proportion of less economical small prey is inverse to the relationship between nestling mass and prey size. These results suggest that the parental behaviour of reed warblers is adjusted by selection to the needs of an average-sized brood. The overall proportion of insect orders was significantly different between the parasitic and host chicks. This result probably reflects the opportunistic foraging habits of the host. The qualitative difference (proportion of insect orders) between host and cuckoo nestling diets is partly a by-product of unequal length distribution of members of different taxonomic groups. The results of this study are consistent with the SSH.     

The relationship between maternal phenotype and offspring quality: do older mothers really produce the best offspring?

Maternal effects are increasingly recognized as important drivers of population dynamics and determinants of evolutionary trajectories. Recently, there has been a proliferation of studies finding or citing a positive relationship between maternal size/age and offspring size or offspring quality. The relationship between maternal phenotype and offspring size is intriguing in that it is unclear why young mothers should produce offspring of inferior quality or fitness. Here we evaluate the underlying evolutionary pressures that may lead to a maternal size/age-offspring size correlation and consider the likelihood that such a correlation results in a positive relationship between the age or size of mothers and the fitness of their offspring. We find that, while there are a number of reasons why selection may favor the production of larger offspring by larger mothers, this change in size is more likely due to associated changes in the maternal phenotype that affect the offspring size-performance relationship. We did not find evidence that the offspring of older females should have intrinsically higher fitness. When we explored this issue theoretically, the only instance in which smaller mothers produce suboptimal offspring sizes is when a (largely unsupported) constraint on maximum offspring size is introduced into the model. It is clear that larger offspring fare better than smaller offspring when reared in the same environment, but this misses a critical point: different environments elicit selection for different optimal sizes of young. We suggest that caution should be exercised when interpreting the outcome of offspring-size experiments when offspring from different mothers are reared in a common environment, because this approach may remove the source of selection (e.g., reproducing in different context) that induced a shift in offspring size in the first place. It has been suggested that fish stocks should be managed to preserve these older age classes because larger mothers produce offspring with a greater chance of survival and subsequent recruitment. Overall, we suggest that, while there are clear and compelling reasons for preserving older females in exploited populations, there is little theoretical justification or evidence that older mothers produce offspring with higher per capita fitness than do younger mothers.     

Loud calls as a mechanism of social coordination in a fission–fusion taxon, the white-bellied spider monkey (Ateles belzebuth)

Spider monkeys (Ateles spp.) live in social groups that exhibit high levels of fission–fusion dynamics, in which group members form subgroups of varying sizes and compositions. Within these fluid societies, how individuals establish contact with dispersed group members with whom they might choose to associate remains unclear. Long-range vocalizations might facilitate interactions between group members and provide a means of social coordination in fission–fusion societies. We evaluated this possibility for one spider monkey vocalization, the loud call, by examining calling behavior, the relationship between loud calls and changes in subgroup size, and the response of individuals to distant calls and playback experiments in a single study group. We found that 82 % of loud calls were emitted within 30 min of a call from a different location, suggesting that individuals frequently emit loud calls in response to the calls of distant group members. Subgroups that emitted loud calls, especially those that responded to distant calls, were much more likely to experience an increase in subgroup size within an hour after calling than those that did not. Animals also approached distant loud calls more than they avoided or ignored these calls. Finally, playbacks of male calls demonstrated that females respond preferentially to the calls of some individuals over others. Taken together, these results provide support for the hypothesis that spider monkey loud calls function to facilitate and initiate interactions between dispersed group members and suggest that vocal signals can play an important role in influencing social interactions in fission–fusion societies.     

Does the choice of climate baseline matter in ecological niche modelling?

Ecological niche models (ENMs) have multiple applications in ecology, evolution and conservation planning. They relate the known locations of a species to characteristics of its environment (usually climate) over its geographical range. Most ENMs are trained using standard 30-year (1961-1990) or 50-year (1951-2000) baselines to represent current climate conditions. Species occurrence records used as input to the models, however, are frequently collected from time periods that differ from those from which the climate is derived. Since climate variability can be significant within and outside baselines, and the distributions of some plants and animals (e.g., annual plants, insects) can adjust to environmental conditions on much shorter time scales, this mismatch between collection records and climatic baselines may affect the utility and accuracy of model outputs. We investigated how the choice of baseline periods influenced modelling efforts, anticipating that climate baselines derived from the same temporal period as the species records would yield improved ENMs. Ten simulated species’ distributions were modelled using an ENM (Maxent) for (a) occurrences and climates within the same temporal period, based on eighteen 10-year baselines within the 20th century and (b) all available samples and climate baselines from 1951-2000 and 1961-1990. Each model was projected onto all the available 10-year climate scenarios and compared to the models trained on the corresponding scenario. We show that temporal mismatches of species occurrences and climate baselines can result in significantly poorer distribution models. Such temporal mismatch may be unavoidable for many studies, but we emphasize here the need to match the time range of samples and climate data whenever possible.     

Does local competition increase the coexistence of species in intransitive networks?

Competitive intransitivity, a situation in which species' competitive ranks cannot be listed in a strict hierarchy, promotes species coexistence through "enemy's enemy indirect facilitation." Theory suggests that intransitivity-mediated coexistence is enhanced when competitive interactions occur at local spatial scales, although this hypothesis has not been thoroughly tested. Here, we use a lattice model to investigate the effect of local vs. global competition on intransitivity-mediated coexistence across a range of species richness values and levels of intransitivity. Our simulations show that local competition can enhance intransitivity-mediated coexistence in the short-term, yet hinder it in the long-term, when compared to global competition. This occurs because local competition slows species disaggregation, allowing weaker competitors to persist longer in the shifting spatial refuges of intransitive networks, enhancing short-term coexistence. Conversely, our simulations show that, in the long-term, local competition traps disaggregated species in unfavorable areas of the competitive arena, where they are excluded by superior competitors. As a result, in the long-term, global intransitive competition allows a greater number of species to coexist than local intransitive competition.     

Active protection of unrelated offspring against parasitoids. A byproduct of self defense?

Natural enemies exert selection pressure on their prey. Predators and parasitoids drive their prey into the evolution of novel traits to cope with this stress. When eggs and juveniles are the target of enemies, defense strategies may rely on adults. However, it is not easy to predict whether adults should actively protect unrelated offspring. Females of the golden egg bug (Phyllomorpha laciniata) mainly oviposit on conspecifics of either sex. Females can also lay eggs on their food plant. Eggs placed on plants suffer from a higher mortality caused by natural enemies than eggs carried by bugs. Females never carry their own eggs and males are seldom related to the eggs they carry. We experimentally explored if conspecifics protect the eggs by studying the behavioral interaction between P. laciniata individuals and the specialist egg parasitoid Gryon bolivari. All bugs exhibited active responses against parasitoids regardless of the sex of the bug, the egg load, and their mating status. Most of the responses prevented parasitoids from reaching the eggs, and thus they reduced the risk of egg parasitization. Although responses of bugs were effective to overcome parasitoid attacks, we suggest that egg protection against parasitoids has evolved as a co-opted trait from a general defense of adult bugs against enemies. In this system, egg defense is not an individual's strategy to protect the offspring, but rather a consequence of the eggs being attached to one's body. It may also explain the low parasitization carried eggs suffer in the wild. The results further highlight the idea of conspecifics as an enemy-free space in P. laciniata.     

Decoration supplementation and male–male competition in the great bowerbird (Ptilonorhynchus nuchalis): a test of the social control hypothesis

Many animals use signals to communicate their social status to conspecifics, and the social control hypothesis suggests that social interactions maintain the evolutionary stability of status signals: low-quality individuals signal at a low level to prevent high-quality individuals from “punishing” them. I examined whether the numbers of decorations at bowers are socially controlled in the great bowerbird (Ptilonorhynchus nuchalis). In two populations, I supplemented males with decorations to determine whether they (a) rejected supplemental decorations and (b) experienced increased bower destruction from rivals. In contrast to the social control hypothesis, males in both populations accepted most supplemental decorations. Though the mean destruction rate did not increase during supplementation in either population, one of the study populations (Townsville) exhibited a negative correlation between the numbers of decorations naturally displayed at bowers and the change in destruction rate during the experiment. Townsville males that naturally had few decorations at their bowers also had more decorations stolen by other males during supplementation than males that naturally had many decorations. These results suggest that the numbers of decorations at bowers are an honest signal of the male's ability to defend his display site from rivals in at least one population of the great bowerbird (Townsville), but they do not support the social control hypothesis because males at both sites failed to limit signal expression. I discuss how the external nature of bower decorations and their availability in the environment may influence the costs and benefits of decoration theft and social control.     

Hunted hunters? Effect of group size on predation risk and growth in the Australian subsocial crab spider Diaea ergandros

A reduced predation risk is considered to be a major adaptive advantage of sociality. While most studies are concerned with non-predatory prey species, group-living predators are likely to face similar threats from higher-order predators. We studied the relationship between group size and predation risk in the subsocial crab spider Diaea ergandros by testing predictions from theoretical models including attack abatement as well as the formation of protective retreats. In a field survey, we found predatory clubionid spiders in 35 % of the D. ergandros nests and as predicted, nest size did not correlate with predator presence. In a subsequent laboratory experiment, we observed survival probability, nest construction activity and feeding behaviour including weight development between groups of different sizes as well as in the absence or presence of a predator. Large groups had an advantage in terms of survival and growth compared to smaller groups or single individuals. They also built significantly larger nests than smaller groups, supporting the idea of protective retreat formation being an adaptive benefit to group living. Even though clubionids did attack D. ergandros, they did not significantly affect overall mortality of D. ergandros. The feeding experiment showed that spiders fed on a larger proportion of flies in the presence of a predator. However, these groups gained significantly less weight compared to the control groups, indicating that the potential predators not only act as predators but also as food competitors, constituting a twofold cost for D. ergandros.     

How unpredictable is the individual scanning process in socially foraging mammals?

In group-forming prey species, theory assumes that individuals within groups should scan independently of one another, with vigilance sequences being relatively unpredictable, making interscan durations highly variable. We attempted to detect any divergence from randomness in the scanning process in three mammalian prey species phylogenetically and geographically separated and exposed to different levels of predation: waterbuck, Kobus ellipsiprymnus defassa, under a high observed predation risk, eastern grey kangaroo, Macropus giganteus, still experiencing occasional predation and European roe deer, Capreolus capreolus, under a very low natural predation risk. Our results revealed that the focal interscan duration increased when the duration of the preceding interscan increased, whatever the studied species and the predation risk that its individuals experienced, and decreased with the preceding scan duration in two species under, respectively, occasional and low predation risks. The exponential distribution was the tested model that fitted the observed distributions of interscan durations least well. We discuss what can trigger non-randomness in scanning, through a non-homogenous Poisson process, at both intra-individual and inter-individual levels, particularly with regard to previous studies that have demonstrated synchronisation of vigilance in such mammals. Our results suggest the need to reconsider any assumption of randomness in scanning in the basic model predicting form and frequency of scanning behaviour by prey species.     

Does the spawning of Calanus finmarchicus in high latitudes follow a reproducible pattern?

In the spring of 1989, an experimental study of the spawning behaviour of Calanus finmarchicus was carried out in Malangen, northern Norway. Here, a single cohort of females reproduce from mid-March to May, approximately coinciding with the wax and wane of the spring phytoplankton bloom. An evaluation of population characteristics such as the proportion of adults, sex ratio, as well as gonad maturation and daily productivity of the females clearly reveals three phases within the population's reproductive period. In between incline and decline, the highest spawning rates (on average >20 eggs female^-1 d^-1, equivalent to 5.7% body C d^-1) occur after the males have disappeared from the population and almost all females have mature gonads. During this period, the ratio of adults to copepodid Stage Vs changes from dominance of adults to that of CVs. Although first egg production was observed prior to the phytoplankton increase, it is suggested that the onset of the phytoplankton spring bloom in the first few days of April enhances the final maturation of ovaries in the females and therefore triggers the onset of the main spawning period. The clutch sizes (max. 95 eggs clutch^-1) vary with the age of the females, while the spawning frequencies depend on the available food quantities. The overlap of an estimated minimal 4 wk spawning period for the individuals leads to a main reproductive phase for the population of ca. 3 wk, during which time mean clutch sizes and spawning frequencies are maximal (highest average clutch size: 70 eggs female^-1 clutch^-1, 100 to 60% of the females spawning). This period ends before the end of the phytoplankton bloom. Calculated by stepwise interpolation and summation of the mean daily egg production in the population, an average female produced ca. 600 eggs during the spring bloom in Malangen 1989. We suggest that reproduction and population development of C. finmarchicus in spring follows a reproducible pattern for a given temperature regime and non-limiting food conditions. In the case of clearly identifiable cohorts, it seems possible to trace the state of reproduction by evaluating population parameters.     

Risk Premiums for Environmental Liability: Does Superfund Increase the Cost of Capital?

Superfund liability may impose financial risk on investors and thereby increase firms' costs of capital. We analyze monthly stock returns for 73 chemical companies using several measures of Superfund exposure. Additional exposure appears to increase costs of capital for larger firms, but perhaps not for smaller firms. From 1988 to 1992, we estimate an average increase in cost of capital for 23 larger firms of between 0.25 to 0.40 percentage points per year. The social cost of Superfund-related financial risk in the chemical industry may be as high as $800 million annually or enough to clean up about 20 sites.     

Does temporal and spatial segregation explain the complex population structure of humpback whales on the coast of West Africa?

Humpback whales (Megaptera novaeangliae) in the Southeastern Atlantic Ocean (International Whaling Commission ‘Breeding Stock B’—BSB) are distributed from the Gulf of Guinea to Western South Africa. Genetic data suggest that this stock may be sub-structured, but it remains unknown if this is due to reproductive segregation. This paper evaluates the spatial and temporal population structure of BSB humpback whales using a combination of maternally and bi-parentally inherited markers. The genetic differentiation that we identify in this study could be due to a combination of (1) spatial and/or temporal segregation on breeding grounds in the greater Gulf of Guinea, (2) the possibility of maternally inherited site fidelity to specific feeding grounds and (3) the use of two generalized but exclusive migratory routes (coastal and offshore) between feeding and breeding areas. Further, photo-identification and genetic sampling efforts in other areas of the Sub-Saharan Western Africa winter range and targeted deployment of satellite tags would help to clarify some of the apparent complexity in the population structure of animals biopsied in this region.     

Should the federal government subsidize industrial pollution control investment?

Tax-exempt financing of industrial pollution control equipment has increased rapidly in recent months. This method of subsidizing pollution abatement is known to be highly inefficient. If tax-exempt industrial revenue bonds were replaced by a more efficient subsidy the net financial benefits to polluting industries could probably be about doubled without any extra cost to American taxpayers. Increased utilization of an inefficient subsidy device and the international energy crisis make this a propitious time for tax experts and economists to re-examine the question, should industrial pollution control investments be subsidized?     

Does the environmental inequality matter? A literature review

Environmental Geochemistry and Health - The environmental inequality theory reveals that the risk of environmental pollution exposure varies among regions and groups and that particular groups face...     

Is sexual monomorphism a predictor of polygynandry? Evidence from a social mammal, the collared peccary

Sexual dimorphism is common in polygynous species, and there is clear evidence that both intra-sexual competition and female preferences can drive the evolution of large body size in males. In contrast, sexual monomorphism is often argued to reflect a relaxation of male mate competition or an intensification of resource competition among females. Alternatively, it might imply opportunities for females to circumvent or counteract male mate competition in a polygynandrous mating system. We test the prediction that sexual monorphism is associated with polygynandry in the collared peccary (Pecari tajacu, Tayassuidae), a social ungulate closely related to the old-world suids. The genetic mating system in the Tayassuidae is unknown, but its sexual monomorphism presents a striking contrast to the strong size dimorphism found in most Suidae, so that a departure from the polygynous system common in Suidae would be noteworthy. We characterized genetic relationships among adults within herds in three geographically distinct populations, assigned parents to 75 offspring, and tested for skew in individual reproductive success. Parentage assignment data indicated that multiple males sire offspring within a herd, and in the population for which genetic data were most complete, 19% of parentage assignments were potentially sired by extra-herd males. Some litters have multiple sires, and neither males nor females monopolized reproduction, even in small herds. This result supports our prediction and suggests that sexual monomorphism may either select for or be an evolutionary consequence of a promiscuous mating system.     

Does phylogenetic inertia explain the evolution of ineffective antipredator behavior in a sunfish-salamander system?

The streamside salamander, Ambystoma barbouri, exhibits ineffective antipredator behavior (high emergence rate from refuge, and high activity while out of refuge) and thus suffers heavy predation in stream pools with sunfish. A. barbouri evolved relatively recently from an ancestor that closely resembled a sister species, A. texanum, which breeds in fishless, ephemeral ponds. Sunfish thus represent a relatively new selection pressure for A. barbouri. Phylogenetic inertia predicts that (1) A. texanum should be very poor at coping with fish and (2) because it has only recently been exposed to fish, A. barbouri should still be poor at avoiding fish, but due to its recent exposure to fish, A. barbouri should be better than A. texanum at coping with sunfish. Experimental results provided mixed support for these predictions. As predicted, A. texanum suffered heavy sunfish predation. Compared to A. texanum, A. barbouri showed a greater tendency to initiate alarm moves that enhanced escape success from fish. However, in both the presence and absence of fish, A. barbouri showed higher emergence rates from refuge and higher movement while out of refuge than A. texanum. These behaviors tend to increase exposure to sunfish, i.e., for these key behaviors, A. barbouri apparently evolved in the wrong direction as far as fish predation is concerned. Due to these offsetting effects (increased exposure to fish, increased escape success), A. barbouri is no better at surviving with sunfish than A. texanum. A possible explanation for the high activity of A. barbouri is its use of highly ephemeral habitats (relative to A. texanum) that favor the evolution of higher activity, feeding, and developmental rates for A. barbouri relative to A. texanum. Received: 15 May 2000 / Revised: 3 August 2000 / Accepted: 18 August 2000